Temporal variations in the reproductive condition of Drosophila obscura Fallén and D. subobscura collin

Summary The reproductive condition, insemination status and fat content of Drosophila obscura and D. subobscura were investigated in samples from natural populations, and in near-natural and artificial experimental situations. In Northern England D. obscura populations pass through either three of four generations per year, one of which includes a period of adult diapause which involves both reproductive condition and fat content. Appropriate day lengths lead to both the initiation and the breaking of this diapause. D. subobscura populations develop continuously and pass through from four to six generations per year. Females from both species were only rarely fertilised when immature. The dependence on both season and food-regime of development, rates in the various stages of both species have been examined and discussed.     

Is there reproductive diapause in the winterform of pear psylla Cacopsyla bidens?

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The distribution and abundance of the British fungal-breeding Drosophila

Abstract.

• 1 The distribution and abundance of five Drosophila species breeding in fungi are examined by attracting adult flies to baited traps and collecting fungal fruit bodies containing the larval stages.
• 2 Changes in species frequency follow a temporal (seasonal and yearly), rather than a spatial (between and within woods) pattern.
• 3 There is a regular diurnal pattern of activity in D.subobscura but not in D.phalerata. Temperature, light intensity, humidity and wind speed have no effect upon the daily activity of D.phalerata. Only temperature affects the activity of D.subobscura.
• 4 Out of 125 species of fungi collected, only forty-eight species produced Drosophila. However, the flies are even more selective, since only five species of fungi (Phallus impudicus, Polyporus squamosus, Amanita rubescens, Pluteus cervinus and Lactarius quietus) produce 80% of all Drosophila reared.
• 5 The commonest species, D.phalerata, has two or three generations per year with a winter diapause. An outline of the yearly life cycle is given, with the major breeding sites for each generation indicated.
• 6 Measures of niche overlap calculated from the data on individual fruit bodies are very low, suggesting that the five species are effectively isolated during the larval period. This ecological isolation is achieved by a summation of several niche dimensions, woodlands, seasons, fungal species, and individual fruiting body differences.

     

An ecological classification of European Drosophila species

Summary The associations shown between species of Drosophila collected in three European countries are analysed using a clustering method. The resulting dendrograms are combined to give a plan of associations shown by all three surveys. These general groupings are interpreted in the light of what is known about Drosophila breeding sites.One ecological group, the fungal breeding species are examined in detail and their pattern of geographical associations investigated. The three most abundant species in collections, D. transversa, D. phalerata and D. cameraria appear to replace one another in a north-south direction in western Europe. It is suggested that ecologically marginal areas may be defined using the frequency of a species within its ecological group.     

Host-plant mediated geographic variation in the life history of Platycotis vittata (Homoptera: Membracidae)

Abstract.

• 1 The oak treehopper Platycotis vittata (Fabricius) is bivoltine throughout its range with discrete generations in the early spring and autumn. Females of both generations diapause.
• 2 Latitudinal variation in the timing of life history exists among four populations of P. vittata along the eastern coastal plain of the United States.
• 3 We disrupted synchronization between P.vittata life history and host-plant phenology in the springs of 1984 and 1985 by placing southern populations on northern trees with a different phenology. This caused some females to oviposit abnormally in late spring, without entering diapause. The offspring of these females had reduced reproductive success.
• 4 Winter diapause can be broken by providing females with a newly flushing tree with high levels of amino nitrogen in the sap. Females terminated winter diapause on trees breaking winter dormancy while females on trees still in winter dormancy did not, under identical photoperiod and temperature regimes.
• 5 The role of host-plant mediated life history variation as a process promoting intraspecific geographic differentiation in the arboreal sap-feeding guild is discussed.

     

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Differences in the pre‐diapause and pre‐oviposition accumulation of critical nutrients in adult females of the beetle Colaphellus bowringi

In many insect species, the differentiation of development between diapause and reproduction first becomes obvious during the diapause preparation (pre‐diapause) and pre‐oviposition phases. However, the differentiation of nutrient accumulation between these two phases remains unclear. We compared the weights of pre‐diapause and reproductive adult female Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), and measured their triacylglycerol (TAG), protein, and carbohydrate content from emergence until they had fed for 4 days post‐eclosion. We also compared the ovarian development and accumulation of lipid droplets between pre‐diapause and reproductive adult females in order to determine whether we could visually detect differences in nutrient allocation. The weights of both pre‐diapause and reproductive females increased with duration of feeding. The fresh weight and water content of pre‐diapause females was significantly lower than that of reproductive females after feeding for 3 days post‐eclosion. Pre‐diapause females channeled their reserves into TAG in the fat body, whereas reproductive females converted nutrients into proteins and carbohydrates for egg development. These results quantify differences in nutrient accumulation between pre‐diapause and reproductive adult female C. bowringi, and provide clues for understanding the molecular mechanisms underlying differences in the allocation of nutrients between diapause and reproduction in insects.     

Differences in male courtship song among the species of thequinaria group ofDrosophila

Courtship song is known to vary among several groups of closely related species ofDrosophila. The present study investigates differences in song among four members of thequinaria group of fungal-breedingDrosophila which are known to coexist in Britain and the near-continent. Results show that the species can be distinguished by several song parameters including interpulse interval.D. phalerata is very different from the other three species, while there is considerable overlap betweenD. kuntzei andD. limbata in several characters.D. transversa appears to be more similar to the latter species than it is toD. phalerata. The question of mate choice and the potential for reproductive character displacement are discussed.     

Feeding,reproductive and locomotor activities in diapausing and non-diapausing adults ofDrosophila

Feeding, reproductive and locomotor activities of fourDrosophila species were studied under short and long daylengths at 15°C. A short daylength induced firm reproductive diapause in experimental strains ofD. subauraria andD. triauraria from northern Japan, but very shallow diapause in those ofD. lutescens andD. rufa from southern Japan. A subtropical strain ofD. triauraria had no diapause. The influence of diapause on feeding activity was detected only in aged (> 12 day old) females; that is, the feeding activity was lower in diapausing females than in non-diapausing ones. Females that do not produce eggs would not require so much energy. On the other hand, young adults of the study species exhibited a high feeding activity and rapidly increased bodyweight irrespective of sex and the diapause state. They would need nutrition to build up their adult body. In males, the feeding activity decreased with age irrespective of the diapause state. Males would not require so much energy for reproductive activity. Diapausing males became heavier than non-diapausing males, perhaps because they accumulated triacylglycerols in fat bodies. However, female bodyweight did not differ by the diapause state, perhaps because diapausing females accumulated triacylglycerols and reproducing females had eggs in their ovaries. InD. triauraria, diapausing individuals exhibited somewhat lower locomotor activity than non-diapausing ones.     

The effect of cold temperature exposure and long-day photoperiod on the termination of the reproductive diapause of newly emerged female Pissodes strobi (Coleoptera: Curculionidae)

Abstract

• 1 There is confusion in the literature concerning a possible reproductive diapause in the adult white pine weevil Pissodes strobi.
• 2 We evaluated the effects of temperature, photoperiod, feeding substrate and mating status on the sexual maturation and oviposition of female white pine weevils.
• 3 Less than 30% of female P. strobi became sexually mature and laid eggs without experiencing dormancy under a temperature regime of 2 °C for 4 weeks.
• 4 Among the females that experienced a cold temperature treatment after emergence, 80% laid eggs after dormancy when exposed to a long‐day (LD 16 : 8 h) photoperiod and 17.6% laid eggs when exposed to a short‐day (LD 8 : 16 h) photoperiod.
• 5 Significantly more eggs were laid by all the females (with and without a cold treatment) when subjected to a long‐day photoperiod compared with a short‐day photoperiod.
• 6 A period of cold temperature followed by exposure to a long‐day photoperiod with warmer temperatures is required to break reproductive diapause and to obtain a good oviposition response in female P. strobi.
• 7 This study reveals the existence of much intraspecific variation in the response of the white pine weevil to temperature and photoperiod with respect to the induction and termination of reproductive diapause.

     

Presence of larval and adult diapauses in a subtropical scarab beetle: graded thermal response for synchronized sexual maturation and reproduction

Abstract The white grub Dasylepida ishigakiensis has a 2‐year life cycle and spends approximately 9 months as a nonfeeding larva, pupa and adult on a subtropical island. Evidence is presented indicating that this beetle has two diapauses that appear to synchronize this long life cycle with the seasons. Larvae exposed to 20, 22.5, 25 and 27.5 °C late in the third (last) stadium pupate rapidly except for some individuals kept at the highest temperature. The latter pupate upon transfer to 22.5 °C, indicating that larval diapause is maintained at high temperature but terminates upon transfer to a lower temperature. Pupal development is directly temperature‐dependent in the range 20–30 °C. Adults develop reproductive organs (i.e. the ovary in females and the seminal vesicles and accessory glands in males) rapidly at 15 and 20 °C, whereas those kept at 25 °C take a long time to do so. Ovarian development is completely suppressed at 30 °C but initiated upon transfer to 20 °C. In the laboratory, males with well‐developed reproductive organs mate even with sexually immature females , whereas females with undeveloped ovaries show no sexual behaviour. Although the two diapauses of this species are thermally regulated (i.e. a characteristic commonly expressed by insects in summer diapause), adults of this beetle emerge from pupae late in the autumn and remain in the soil for 2 months. Adult diapause effectively serves to synchronize the time of sexual maturation with the coldest month of the year.     

Effects of diapause on post‐diapause reproductive investment in the moth Ostrinia scapulalis

Diapause is a strategy used by many insect species to survive adverse environmental conditions. However, diapause incurs costs that may have adverse effects on post‐diapause development and reproduction. We herein investigated the effects of diapause on the post‐diapause reproductive investment of males and females in a multivoltine moth, the adzuki bean borer, Ostrinia scapulalis (Walker) (Lepidoptera: Crambidae). We found that (1) post‐diapause males and females were smaller and had lower mating success than non‐diapause individuals, (2) post‐diapause females had lower fecundity and shorter longevity than non‐diapause females, (3) post‐diapause males transferred similar numbers of eupyrene and apyrene sperm as non‐diapause males, (4) the fecundity and longevity of non‐diapause females mated with post‐diapause males and those mated with non‐diapause males were not significantly different, and (5) no significant relationship was found between diapause duration (short and long) and post‐diapause reproductive investments in both males and females. These results suggest that post‐diapause males did not reduce reproductive investment in spite of the cost of diapause, and the significant decline in reproductive output in post‐diapause females was due to their reduced body weight and longevity, which appeared to be direct consequences of the cost of diapause.     

Environmental Cues,Endocrine Factors,and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.     

Time of the season: the effect of host photoperiodism on diapause induction in an insect herbivore,Leptinotarsa decemlineata

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Nocturnal activity and resource utilization in the aphid hyperparasitoid, Dendrocerus carpenteri

Abstract.

• 1 Females of the aphid hyperparasitoid Dendrocerus carpenteri (Curtis) search successfully for hosts during both day and night. Oviposition numbers per host patch did not differ significantly between day and night.
• 2 D.carpenteri females also displayed a nocturnal flight activity, showing that they are not only capable of searching on a given host plant but also of dispersing between host plants.
• 3 Nocturnal oviposition activity was mainly influenced by egg load. Females with a high egg load laid more eggs at night than females with a comparatively low egg load. Thus, D.carpenteri females may use nocturnal foraging to compensate for the lack of oviposition opportunities during day.
• 4 D.carpenteri females which foraged continuously for hosts both day and night (= for 24 h per day) benefitted from an 1.4-fold increase in lifetime reproductive success when compared to females which foraged only by day (= for 16 h per day).
• 5 The benefit of night foraging for this species is a significantly increased reproductive success.

     

Variation in diapause and sex ratio in the parasitoidAsobara tabida

Asobara tabida Nees (Hymenoptera: Braconidae) is a widespread parasitoid, attacking larvae ofDrosophila (Diptera: Drosophilidae) species in fermenting substrates. In this species, geographic variation is found in the percentage of parasitoids entering diapause and in the sex ratio of emerging parasitoids. Percentage diapause appears to be influenced by host species (more parasitoids enter diapause inD. melanogaster Meigen than inD. subobscura Collin) and temperature. It is not correlated with any of the abiotic factors investigated, but is correlated with survival probability inD. melanogaster larvae and with the time of year in which the experiment was conducted (even though none of the parasitoids experienced natural day light). Sex ratio was only found to correlate with percentage diapause, suggesting that males enter diapause more frequently than females. It is concluded thatA. tabida uses diapause to survive both unfavourable abiotic and biotic circumstances.     

Biennial reproduction with embryonic diapause in Lopholithodes foraminatus (Anomura: Lithodidae) from British Columbia waters

Abstract. A paucity of data on the reproductive cycle of crabs in the family Lithodidae inhibits both the development of management strategies and the formulation of hypotheses regarding the evolution of lithodid life histories. Life‐history parameters of Lopholithodes foraminatus from British Columbia, Canada, were investigated based on 26 females maintained in the laboratory and supplementary observations on other living and preserved animals. The rate of embryonic development was determined by measuring the percentage area occupied by yolk in lateral views of eggs removed from brooding females throughout development. Females of L. foraminatus exhibited biennial reproduction including an 18‐month brooding period. Females molted, mated, and extruded eggs in mid‐summer, and did not release larvae until late winter or early spring of the second year after fertilization. Embryogenesis included a 12‐month diapause at the gastrula stage. Females released larvae over a mean interval of 69 d, the longest reported for any lithodid. While the development stage of embryos was observed to be heterogeneous within a brood, no spatial gradient in development rate was observed, calling into question the oxygen limitation hypothesis of extended hatching. Biennial reproduction of individuals of L. foraminatus may be a consequence of a relatively low‐quality habitat. Relative to annual reproduction, biennial reproduction halves the potential rate of increase of a population and increases vulnerability to overharvesting, suggesting that L. foraminatus is not a good candidate for commercial exploitation. The adaptive value of embryonic diapause is uncertain and warrants further research.     

Effects of nutrition and methoprene treatment upon reproductive diapause in Caloptilia fraxinella (Lepidoptera: Gracillariidae)

Abstract Female Caloptilia fraxinella exhibit a prolonged reproductive diapause immediately post adult emergence in mid‐summer until the next spring when mating, egg development and oviposition on fresh Fraxinus spp. leaflets occur. Factors that effect the termination of reproductive diapause are investigated in this species. Caloptilia fraxinella diapausing adults held in overwintering conditions (2 °C, LD 0 : 24 h) for 24 weeks terminate diapause after placement for 2 weeks in simulated summer conditions (24 °C, LD 16 : 8 h) only if they are provided with 10% sugar water. Exogenous application of the Juvenile Hormone (JH) analogue methoprene to moths in both early‐ (summer) and mid‐ (autumn) reproductive diapause demonstrates that JH affects diapause termination but a carbohydrate nutrition source also mediates mating and vitellogenesis. Mating between moth pairs early in diapause occurs only after treatment with methoprene and provision with sugar water. However, there is no impact of mating on the propensity of females to produce vitellogenic oöcytes. Moths collected in the autumn in mid‐diapause respond in a dose‐dependent fashion to methoprene treatment and the response is greater than that of moths early in diapause collected in the summer. Treatment with methoprene and access to sugar water results in vitellogenic oöcytes in 18.75% of females from mid‐diapause moth pairs treated with 0.01 μg methoprene per insect and in all females from pairs treated at the two highest doses of methoprene (0.1 and 1 μg per insect). Mating occurs only between moths in mid‐diapause treated with the two highest doses of methoprene and these doses induce 91% and 100% mating, respectively. Both control and methoprene‐treated males in mid‐diapause held under summer conditions mate successfully and pass a spermatophore to their methoprene‐treated female partner. These data demonstrate that female C. fraxinella undergo a prolonged reproductive diapause in which termination is dependent on JH and further mediated by a carbohydrate nutrition source. The production of vitellogenic oöcytes is independent of mating. These data also provide evidence that response of moths in diapause to exogenous applications of methoprene differs throughout the diapause period and between male and female C. fraxinella.     

Egg size increases with maternal age in the cotton stainer bugs Dysdercus fasciatus and D.cardinalis (Hemiptera: Pyrrhocoridae)

Abstract.

• 1 Egg-size variation over the reproductive span of laboratory-reared females is described in two species of cotton stainers: Dysdercus fasciatus Sign. from woody Malvales and D.cardinalis Gerst. from herbaceous Malvales.
• 2 Egg size increases with maternal age due to a decrease in clutch size as maternal age advances in both species.
• 3 The two species are similar in size, but egg size is about 28% larger and clutch size about 19% smaller in D.fasciatus than in D.cardinalis. These contrasts may be related to the host-plant biology which differs between the two species.

     

The effect of larval photoperiod on pupal colour and diapause in swallowtail butterflies

ABSTRACT.

• 1 There are significant differences in the effects of larval photo-period on diapause and pupal colour among the species Papilio polyxenes Fabr., P.troilus L., Battus philenor (L.) and Eurytides marcellus (Cramer).
• 2 Diapause and pupal colour in P.polyxenes and P.troilus are strongly influenced by larval photoperiod, short photophase eliciting brown diapausing pupae. Photoperiods of 15L:9D permit the expression of the green and brown pupal colour alternatives.
• 3 Pupal colour in B.philenor and E.marcellus is not affected by larval photoperiod, but short photophase induces diapause in these species.
• 4 All species except B.philenor show an association between brown pupal colour and diapause: Emarcellus when reared on long (midsummer) photophase, P.polyxenes and P.troilus when reared on short (autumnal) photophase.
• 5 In P.polyxenes, short photophase can affect pupal colour responses directly, whether the individual enters diapause or not.
• 6 Differences among the species are related to differences in the ecology of their natural pupation sites.